Onur Güntürkün

Ruhr-Universität Bochum, Bochum, North Rhine-Westphalia, Germany

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Publications (262)1027.21 Total impact

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    Frontiers in Psychology 09/2014; · 2.80 Impact Factor
  • Onur Güntürkün, Maik C. Stüttgen, Martina Manns
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    ABSTRACT: Deeper understanding of the neuronal basis of behavior and its evolution requires investigation of model organisms taken from different taxonomic groups. The merits of this comparative approach are highlighted by research on birds: while their cognitive capacities have long been underestimated, research on avian model systems more recently has begun to provide central insights into the functional organization of the brain. In particular, domesticated homing pigeons (Columba livia) have been used as a model for the study of the psychological processes underlying learning, memory, and choice behavior, and much of current animal learning theory is based on findings with pigeons. Moreover, the vast amount of available behavioral and anatomical data has rendered the pigeon one of the key model species of behavioral and comparative neuroscience. This article illustrates some insights gained from research with pigeons with applicability beyond the class of aves.
    Neuroforum 09/2014; · 0.04 Impact Factor
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    ABSTRACT: Sensory systems provide organisms with information on the current status of the environment, thus enabling adaptive behavior. The neural mechanisms by which sensory information is exploited for action selection are typically studied with mammalian subjects performing perceptual decision-making tasks, and most of what is known about these mechanisms at the single-neuron level is derived from cortical recordings in behaving monkeys. To explore the generality of neural mechanisms underlying perceptual decision making across species, we recorded single-neuron activity in the pigeon nidopallium caudolaterale (NCL), a non-laminated associative forebrain structure thought to be functionally equivalent to mammalian prefrontal cortex, while subjects performed a visual categorisation task. We found that, whereas the majority of NCL neurons unspecifically upregulated or downregulated activity during stimulus presentation, ~20% of neurons exhibited differential activity for the sample stimuli and predicted upcoming choices. Moreover, neural activity in these neurons was ramping up during stimulus presentation and remained elevated until a choice was initiated, a response pattern similar to that found in monkey prefrontal and parietal cortices in saccadic choice tasks. In addition, many NCL neurons coded for movement direction during choice execution and differentiated between choice outcomes (reward and punishment). Taken together, our results implicate the NCL in the selection and execution of operant responses, an interpretation resonating well with the results of previous lesion studies. The resemblance of the response patterns of NCL neurons to those observed in mammalian cortex suggests that, despite differing neural architectures, mechanisms for perceptual decision making are similar across classes of vertebrates.
    European Journal of Neuroscience 08/2014; · 3.75 Impact Factor
  • Sarah Starosta, Maik C Stüttgen, Onur Güntürkün
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    ABSTRACT: While the subject of learning has attracted immense interest from both behavioral and neural scientists, only relatively few investigators have observed single-neuron activity while animals are acquiring an operantly conditioned response, or when that response is extinguished. But even in these cases, observation periods usually encompass only a single stage of learning, i.e. acquisition or extinction, but not both (exceptions include protocols employing reversal learning; see Bingman et al.(1) for an example). However, acquisition and extinction entail different learning mechanisms and are therefore expected to be accompanied by different types and/or loci of neural plasticity. Accordingly, we developed a behavioral paradigm which institutes three stages of learning in a single behavioral session and which is well suited for the simultaneous recording of single neurons' action potentials. Animals are trained on a single-interval forced choice task which requires mapping each of two possible choice responses to the presentation of different novel visual stimuli (acquisition). After having reached a predefined performance criterion, one of the two choice responses is no longer reinforced (extinction). Following a certain decrement in performance level, correct responses are reinforced again (reacquisition). By using a new set of stimuli in every session, animals can undergo the acquisition-extinction-reacquisition process repeatedly. Because all three stages of learning occur in a single behavioral session, the paradigm is ideal for the simultaneous observation of the spiking output of multiple single neurons. We use pigeons as model systems, but the task can easily be adapted to any other species capable of conditioned discrimination learning.
    Journal of Visualized Experiments 06/2014; 88.
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    Emre Unver, Onur Güntürkün
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    ABSTRACT: In birds each hemisphere receives visual input from the contralateral eye. Since birds have no corpus callosum, avian brains are often seen as 'natural split brains'. How do birds cope with situations, when both hemispheres are brought into conflict? If under such conditions one hemisphere completely determines the response, this is called meta-control. This phenomenon has recently been demonstrated in pigeons. The aim of the current study is to test, if meta-control results from an interhemispheric conflict that would require interhemispheric interaction, possibly via the commissura anterior. To this end, we trained pigeons in a forced-choice color discrimination task under monocular condition such that each hemisphere was trained with a different pair of colors. Subsequently, pigeons were binocularly tested with conflicting and non-conflicting stimulus patterns. Conflicting stimuli indeed produced a delayed reaction time as expected when two divergent decisions create a conflict. In addition, we sometimes observed a pecking pattern that seemed to represent the average of two discrepant and hemisphere-specific movements. Thus, pigeons possibly undergo interhemispheric conflict during meta-control even without a corpus callosum. However, also when having decided to peck a certain color, the planned movement trajectory of the other hemisphere sometimes compromises the final pecking movement.
    Behavioural brain research. 05/2014;
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    ABSTRACT: Dominance of the left hemisphere for many aspects of speech production and perception is one of the best known examples of functional hemispheric asymmetries in the human brain. Classic theories about its ontogenesis assume that it is determined by the same ontogenetic factors as handedness because the two traits are correlated to some extent. However, the strength of this correlation depends on the measures used to assess the two traits, and the neurophysiological basis of language lateralization is different from that of handedness. Therefore, we argue that although the two traits show partial pleiotropy, there is also a substantial amount of independent ontogenetic influences for each of them. This view is supported by several recent genetic and neuroscientific studies that are reviewed in the present article.
    Neuroscience & Biobehavioral Reviews 04/2014; · 10.28 Impact Factor
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    ABSTRACT: Severe feather pecking (FP) is a detrimental behavior causing welfare problems in laying hens. Divergent genetic selection for FP in White Leghorns resulted in strong differences in FP incidences between lines. More recently, it was shown that the high FP (HFP) birds have increased locomotor activity as compared to hens of the low FP (LFP) line, but whether these lines differ in central serotonin (5-hydroxytryptamine, 5-HT) release is unknown. We compared baseline release levels of central 5-HT, and the metabolite 5-HIAA in the limbic and prefrontal subcomponents of the caudal nidopallium by in vivo microdialysis in adult HFP and LFP laying hens from the ninth generation of selection. A single subcutaneous d-fenfluramine injection (0.5mg/kg) was given to release neuronal serotonin in order to investigate presynaptic storage capacity. The present study shows that HFP hens had higher baseline levels of 5-HT in the caudal nidopallium as compared to LFP laying hens. Remarkably, no differences in plasma tryptophan levels (precursor of 5-HT) between the lines were observed. D-fenfluramine increased 5-HT levels in both lines similarly indirectly suggesting that presynaptic storage capacity was the same. The present study shows that HFP hens release more 5-HT under baseline conditions in the caudal nidopallium as compared to the LFP birds. This suggests that HFP hens are characterized by a higher tonic 5-HT release.
    Behavioural brain research 04/2014; · 3.22 Impact Factor
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    ABSTRACT: Social communication through touch and mutual grooming can convey highly salient socio-emotional signals and has been shown to involve the neuropeptide oxytocin (OXT) in several species. Less is known about the modulatory influence of OXT on the neural and emotional responses to human interpersonal touch. The present randomized placebo (PLC)-controlled within-subject pharmaco-functional magnetic resonance imaging (fMRI) study was designed to test the hypothesis that a single intranasal dose of synthetic OXT (24 IU) would facilitate both neural and emotional responses to interpersonal touch in a context (female vs male touch) and trait (autistic trait load) specific manner. Specifically, the experimental rationale was to manipulate the reward value of interpersonal touch independent of the intensity and type of actual cutaneous stimulation administered. Thus forty heterosexual males believed they were touched by either a man or a woman, although in fact an identical pattern of touch was always given by the same female experimenter blind to condition type. Our results show that OXT increased the perceived pleasantness of female, but not male touch, and associated neural responses in the insula, precuneus, orbitofrontal and pregenual anterior cingulate cortex. Moreover, the behavioral and neural effects of OXT were negatively correlated with autistic-like traits. Taken together, this is the first study to show that the perceived hedonic value of human heterosexual interpersonal touch is facilitated by OXT in men, but that its behavioral and neural effects in this context are blunted in individuals with autistic traits.Neuropsychopharmacology accepted article peview online, 03 April 2014; doi:10.1038/npp.2014.78.
    Neuropsychopharmacology: official publication of the American College of Neuropsychopharmacology 04/2014; · 8.68 Impact Factor
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    ABSTRACT: The majority of experiments exploring context-dependent extinction learning employ Pavlovian fear conditioning in rodents. Since mechanisms of appetitive and aversive learning are known to differ at the neuronal level, we sought to investigate extinction learning in an appetitive setting. Working with pigeons, we established a within-subject ABA renewal paradigm based on Rescorla (QJ Exp Psychol 61:1793) and combined it with pharmacological interventions during extinction. From the fear conditioning literature, it is known that both prefrontal cortex and the hippocampus are core structures for context-specific extinction learning. Accordingly, we transiently inactivated the nidopallium caudolaterale (NCL, a functional analogue of mammalian prefrontal cortex) and the hippocampus in separate experiments by intracranial infusion of the sodium-channel blocker tetrodotoxin immediately before extinction training. We find that TTX in both structures non-specifically suppresses conditioned responding, as revealed by a reduction of response rate to both the extinguished conditioned stimulus and a control stimulus which remained reinforced throughout the experiment. Furthermore, TTX during extinction training impaired later extinction retrieval assessed under drug-free conditions. This was true when responding to the extinguished stimulus was assessed in the context of extinction but not when tested in the context of acquisition, although both contexts were matched with respect to their history of conditioning. These results indicate that both NCL and hippocampus are involved in extinction learning under appetitive conditions or, more specifically, in the consolidation of extinction memory, and that their contribution to extinction is context-specific.
    Behavioural brain research 02/2014; · 3.22 Impact Factor
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    ABSTRACT: Efforts to understand nervous system structure and function have received new impetus from the federal Brain Research through Advancing Innovative Neurotechnologies (BRAIN) Initiative. Comparative analyses can contribute to this effort by leading to the discovery of general principles of neural circuit design, information processing, and gene-structure-function relationships that are not apparent from studies on single species. We here propose to extend the comparative approach to nervous system ‘maps’ comprising molecular, anatomical, and physiological data. This research will identify which neural features are likely to generalize across species, and which are unlikely to be broadly conserved. It will also suggest causal relationships between genes, development, adult anatomy, physiology, and, ultimately, behavior. These causal hypotheses can then be tested experimentally. Finally, insights from comparative research can inspire and guide technological development. To promote this research agenda, we recommend that teams of investigators coalesce around specific research questions and select a set of ‘reference species’ to anchor their comparative analyses. These reference species should be chosen not just for practical advantages, but also with regard for their phylogenetic position, behavioral repertoire, well-annotated genome, or other strategic reasons. We envision that the nervous systems of these reference species will be mapped in more detail than those of other species. The collected data may range from the molecular to the behavioral, depending on the research question. To integrate across levels of analysis and across species, standards for data collection, annotation, archiving, and distribution must be developed and respected. To that end, it will help to form networks or consortia of researchers and centers for science, technology, and education that focus on organized data collection, distribution, and training. These activities could be supported, at least in part, through existing mechanisms at NSF, NIH, and other agencies. It will also be important to develop new integrated software and database systems for cross-species data analyses. Multidisciplinary efforts to develop such analytical tools should be supported financially. Finally, training opportunities should be created to stimulate multidisciplinary, integrative research into brain structure, function, and evolution. J. Comp. Neurol., 2014. © 2014 Wiley Periodicals, Inc.
    The Journal of Comparative Neurology 02/2014; · 3.66 Impact Factor
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    ABSTRACT: The avian hippocampal formation (HF) and mammalian hippocampus share a similar functional role in spatial cognition, but the underlying neuronal mechanisms enabling the functional similarity are incompletely understood. To better understand the organization of the avian HF and its transmitter receptors, we analyzed binding site densities for glutamatergic AMPA, NMDA and kainate receptors, GABAA receptors, muscarinic M1 , M2 and nicotinic (nACh) acetylcholine receptors, noradrenergic α1 and α2 receptors, serotonergic 5-HT1A receptors and dopaminergic D1/5 receptors using quantitative in vitro receptor autoradiography. Additionally, we performed a modified Timm staining procedure to label zinc. The regionally different receptor densities mapped well on to seven HF subdivisions previously described. Several differences in receptor expression highlighted distinct HF subdivisions. Notable examples include 1) high GABAA and α1 receptor expression, which rendered distinctive ventral subdivisions, 2) high α2 - receptor expression which rendered distinctive a dorsomedial subdivision, 3) distinct kainate, α2 and muscarinic receptor densities that rendered distinctive the two dorsolateral subdivisions, and 4) a dorsomedial region that was characterized by high kainate receptor density. We further observed similarities in receptor binding densities between subdivisions of the avian and mammalian HF. Despite the similarities, we propose that 300 hundred million years of independent evolution has led to a mosaic of similarities and differences in the organization of the avian HF and mammalian hippocampus, and that thinking about the avian HF in terms of the strict organization of the mammalian hippocampus is likely insufficient to understand the HF of birds. J. Comp. Neurol., 2014. © 2014 Wiley Periodicals, Inc.
    The Journal of Comparative Neurology 01/2014; · 3.66 Impact Factor
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    ABSTRACT: The active generation of behavioral variability is thought to be a pivotal element in reinforcement based learning. One example for this principle is song learning in oscine birds. Oscines possess a highly specialized set of brain areas that compose the song system. It is yet unclear how the song system evolved. One important hypothesis assumes a motor origin of the song system, i.e. the song system may have developed from motor pathways that were present in an early ancestor of extant birds. Indeed, in pigeons neural pathways are present that parallel the song system. We examined whether one component of these pathways, a forebrain area termed nidopallium intermedium medialis pars laterale (NIML), is functionally comparable to its putative homologue, the lateral magnocellular nucleus of the anterior nidopallium (LMAN) of the song system. LMAN conveys variability into the motor output during singing; a function crucial for song learning and maintenance. We tested if NIML is likewise associated with the generation of variability. We used a behavioral paradigm in which pigeons had to find hidden target areas on a touch screen to gain food rewards. Alterations in pecking variability would result in changes of performance levels in this search paradigm. We found that pharmacological inactivation of NIML did not reduce pecking variability contrasting increases of song stereotypy observed after LMAN inactivation.
    Behavioural brain research 01/2014; · 3.22 Impact Factor
  • [Show abstract] [Hide abstract]
    ABSTRACT: Dominance of the left hemisphere for many aspects of speech production and perception is one of the best known examples of functional hemispheric asymmetries in the human brain. Classic theories about its ontogenesis assume that it is determined by the same ontogenetic factors as handedness because the two traits are correlated to some extent. However, the strength of this correlation depends on the measures used to assess the two traits, and the neurophysiological basis of language lateralization is different from that of handedness. Therefore, we argue that although the two traits show partial pleiotropy, there is also a substantial amount of independent ontogenetic influences for each of them. This view is supported by several recent genetic and neuroscientific studies that are reviewed in the present article.
    Neuroscience & Biobehavioral Reviews 01/2014; · 10.28 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Efforts to understand nervous system structure and function have received new impetus from the federal Brain Research through Advancing Innovative Neurotechnologies (BRAIN) Initiative. Comparative analyses can contribute to this effort by leading to the discovery of general principles of neural circuit design, information processing, and gene-structure-function relationships that are not apparent from studies on single species. We here propose to extend the comparative approach to nervous system 'maps' comprising molecular, anatomical, and physiological data. This research will identify which neural features are likely to generalize across species, and which are unlikely to be broadly conserved. It will also suggest causal relationships between genes, development, adult anatomy, physiology, and, ultimately, behavior. These causal hypotheses can then be tested experimentally. Finally, insights from comparative research can inspire and guide technological development. To promote this research agenda, we recommend that teams of investigators coalesce around specific research questions and select a set of 'reference species' to anchor their comparative analyses. These reference species should be chosen not just for practical advantages, but also with regard for their phylogenetic position, behavioral repertoire, well-annotated genome, or other strategic reasons. We envision that the nervous systems of these reference species will be mapped in more detail than those of other species. The collected data may range from the molecular to the behavioral, depending on the research question. To integrate across levels of analysis and across species, standards for data collection, annotation, archiving, and distribution must be developed and respected. To that end, it will help to form networks or consortia of researchers and centers for science, technology, and education that focus on organized data collection, distribution, and training. These activities could be supported, at least in part, through existing mechanisms at NSF, NIH, and other agencies. It will also be important to develop new integrated software and database systems for cross-species data analyses. Multidisciplinary efforts to develop such analytical tools should be supported financially. Finally, training opportunities should be created to stimulate multidisciplinary, integrative research into brain structure, function, and evolution.
    Brain Behavior and Evolution 01/2014; 83:1-8. · 2.89 Impact Factor
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    ABSTRACT: The biological mechanisms underlying long-term partner bonds in humans are unclear. The evolutionarily conserved neuropeptide oxytocin (OXT) is associated with the formation of partner bonds in some species via interactions with brain dopamine reward systems. However, whether it plays a similar role in humans has as yet not been established. Here, we report the results of a discovery and a replication study, each involving a double-blind, placebo-controlled, within-subject, pharmaco-functional MRI experiment with 20 heterosexual pair-bonded male volunteers. In both experiments, intranasal OXT treatment (24 IU) made subjects perceive their female partner's face as more attractive compared with unfamiliar women but had no effect on the attractiveness of other familiar women. This enhanced positive partner bias was paralleled by an increased response to partner stimuli compared with unfamiliar women in brain reward regions including the ventral tegmental area and the nucleus accumbens (NAcc). In the left NAcc, OXT even augmented the neural response to the partner compared with a familiar woman, indicating that this finding is partner-bond specific rather than due to familiarity. Taken together, our results suggest that OXT could contribute to romantic bonds in men by enhancing their partner's attractiveness and reward value compared with other women.
    Proceedings of the National Academy of Sciences 11/2013; · 9.81 Impact Factor
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    ABSTRACT: Sex hormones have been reported to dynamically modulate the expression of implicit motives, a concept that has previously been thought to be relatively stable over time. This study investigates to what extent the need for affiliation, power, and achievement, as well as the form of enactment of these needs as measured with the Operant Motive Test (OMT), are affected by cycle-phase dependent sex hormone fluctuations. In addition to measuring the strength of motive expression, the OMT also captures different forms of motive enactment. In an intra-subject design with repeated measures, no evidence for cycle-phase related variation in overall motive scores was found. However, when different forms of motive enactment were considered, an effect of menstrual cycle was observed. The incentive-based inhibition of the power motive was significantly reduced at the time of ovulation, compared to the menstrual and to the mid-luteal phase, in naturally cycling women. In women with relatively stable hormone concentrations (due to using hormonal contraceptives), no significant changes in the form of motive enactment were evident. The results indicate a specific hormonal influence on motive-related cognitive processes that are related to inhibitive processes in behavior control.
    Physiology & Behavior 10/2013; · 3.16 Impact Factor
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    ABSTRACT: Handedness is the single most studied aspect of human brain asymmetries. For long it has been thought to be a monogenic trait that can produce an asymmetrical shift of cerebral mechanisms, thereby producing right handedness. Nevertheless, a single gene explaining a sufficient amount of phenotypic variance has not been identified. The results of several recent studies using advanced molecular genetic techniques suggest that a multifactorial model taking into account both multiple genetic and environmental factors, as well as their interactions, might be better suited to explain the complex processes underlying the ontogenesis of handedness. In this article, we review the new insights into handedness genetics provided by these studies and discuss, how integrating results from genetic and neuroscientific studies might help us to generate more accurate models of the ontogenesis of handedness. Based on these thoughts, we suggest several candidate gene groups (e.g. genes involved in the formation of the corpus callosum, asymmetrically expressed genes or genes involved in the development of structural left-right asymmetries) whose investigation would help to further understand the complex relation of genes, the brain and handedness.
    Neuroscience & Biobehavioral Reviews 10/2013; · 10.28 Impact Factor
  • Source
    [Show abstract] [Hide abstract]
    ABSTRACT: Handedness is the single most studied aspect of human brain asymmetries. For long it has been thought to be a monogenic trait that can produce an asymmetrical shift of cerebral mechanisms, thereby producing right handedness. Nevertheless, a single gene explaining a sufficient amount of phenotypic variance has not been identified. The results of several recent studies using advanced molecular genetic techniques suggest that a multifactorial model taking into account both multiple genetic and environmental factors, as well as their interactions, might be better suited to explain the complex processes underlying the ontogenesis of handedness. In this article, we review the new insights into handedness genetics provided by these studies and discuss, how integrating results from genetic and neuroscientific studies might help us to generate more accurate models of the ontogenesis of handedness. Based on these thoughts, we suggest several candidate gene groups (e.g. genes involved in the formation of the corpus callosum, asymmetrically expressed genes or genes involved in the development of structural left-right asymmetries) whose investigation would help to further understand the complex relation of genes, the brain and handedness.
    Neuroscience & Biobehavioral Reviews 09/2013; · 10.28 Impact Factor
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    ABSTRACT: Feather pecking (FP) in laying hens may cause mortality due to cannibalism. Novel breeding methods using survival days of group-housed siblings allow for the genetic selection of laying hens with low mortality (LML: low mortality line) due to cannibalism. Previous studies have demonstrated less fear-related behavior and also less FP in LML hens compared to CL. Selection also caused changes in locomotor behavior in an open field. It is unknown, however, whether selection for low mortality affects central neurotransmitter levels. In this study, brain monoamine levels were measured in the dorsal thalamus, medial striatum, hippocampus and arcopallium of adult laying hens of both LML and CL using HPLC. Brain samples were collected after 5-min of manual restraint. The most prominent line differences were found in the arcopallium. Compared to CL, LML had lower levels of noradrenaline (NA) and 3,4-dihydroxyphenylacetic acid (DOPAC) and tended to have lower levels of dopamine (DA), homovanillic acid (HVA), and 5-hydroxyindoleacetic acid (5-HIAA). Levels of serotonin (5-HT), 5-HT- and DA-turnover in this brain area were not affected by line. LML showed less fear-related behavior during the restraint than CL. These findings show that selection for low mortality in hens leads to changes of predominantly the dopaminergic system in the chicken's arcopallium, a forebrain somatomotor area also related to fear. This suggests a relationship between catecholamine functioning in this brain area and FP and cannibalistic behavior in chickens and underpins previously found relationships between FP, fear and high activity.
    Behavioural brain research 09/2013; · 3.22 Impact Factor
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    ABSTRACT: Left-hemispheric language dominance is a well-known characteristic of the human language system, but the molecular mechanisms underlying this crucial feature of vocal communication are still far from being understood. The forkhead box P2 gene FOXP2, which has been related to speech development, constitutes an interesting candidate gene in this regard. Therefore, the present study was aimed at investigating effects of variation in FOXP2 on individual language dominance. To this end, we used a dichotic listening and a visual half-field task in a sample of 456 healthy adults. The FOXP2 SNPs rs2396753 and rs12533005 were found to be significantly associated with the distribution of correct answers on the dichotic listening task. These results show that variation in FOXP2 may contribute to the inter-individual variability in hemispheric asymmetries for speech perception.
    Brain and Language 08/2013; 126(3):279-284. · 3.39 Impact Factor

Publication Stats

6k Citations
1,027.21 Total Impact Points

Institutions

  • 1987–2014
    • Ruhr-Universität Bochum
      • • Biopsychology
      • • International Graduate School of Neuroscience
      • • Abteilung für Cytologie
      • • Fakultät für Psychologie
      Bochum, North Rhine-Westphalia, Germany
  • 2013
    • Imperial College London
      • Department of Computing
      London, ENG, United Kingdom
  • 2012–2013
    • University of Bonn
      • Department of Neurobiology
      Bonn, North Rhine-Westphalia, Germany
  • 2011
    • University of Wuerzburg
      • Department of Neurology
      Würzburg, Bavaria, Germany
  • 2002–2011
    • Goethe-Universität Frankfurt am Main
      • • Neurologisches Institut - Edinger Institut
      • • Institute of Psychology
      Frankfurt am Main, Hesse, Germany
    • Philipps University of Marburg
      Marburg, Hesse, Germany
  • 2006–2010
    • Heinrich-Heine-Universität Düsseldorf
      • • C. u. O. Vogt-Institut für Hirnforschung
      • • Institut für Experimentelle Psychologie
      Düsseldorf, North Rhine-Westphalia, Germany
    • University of Amsterdam
      • Faculty of Science
      Amsterdam, North Holland, Netherlands
  • 2008–2009
    • Queen's University
      • Department of Psychology
      Kingston, Ontario, Canada
  • 2004
    • Marmara University
      • Department of Child and Adolescent Psychiatry
      İstanbul, Istanbul, Turkey
  • 1989–2003
    • Universität Konstanz
      • Faculty of Sciences
      Konstanz, Baden-Wuerttemberg, Germany
  • 1998
    • Universität Basel
      • Botanical Institute
      Basel, BS, Switzerland
  • 1995
    • University of Münster
      Muenster, North Rhine-Westphalia, Germany
  • 1993
    • Rutgers, The State University of New Jersey
      • Center for Molecular and Behavioral Neuroscience
      New Brunswick, NJ, United States
    • University of Maryland, College Park
      • Department of Psychology
      College Park, MD, United States
  • 1992
    • University of Alaska Fairbanks
      • Institute of Arctic Biology
      Fairbanks, AK, United States